Article(id=1304388058355364425, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304388047747969563, articleNumber=null, orderNo=null, doi=10.7501/j.issn.0253-2670.2026.10.021, pmid=null, cstr=null, oa=null, hot=0, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=null, receivedDate=1762012800000, receivedDateStr=2025-11-02, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1788919947470, onlineDateStr=2026-09-09, pubDate=null, pubDateStr=null, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1788919947470, onlineIssueDateStr=2026-09-09, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1788919947470, creator=13701087609, updateTime=1788919947470, updator=13701087609, issue=Issue{id=1304388047747969563, tenantId=1146029695717560320, journalId=1302319053441957962, year='2026', volume='57', issue='10', pageStart='3685', pageEnd='4088', issueExtLink='null', onlineDate='null', pubDate='1779897600000', pubDateStr='2026-05-28', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1788919944940, creator='13701087609', updateTime=1788923403989, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1304402556332037104, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304388047747969563, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1304402556332037105, tenantId=1146029695717560320, journalId=1302319053441957962, issueId=1304388047747969563, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=3926, endPage=3939, ext={EN=ArticleExt(id=1304388060230218315, articleId=1304388058355364425, tenantId=1146029695717560320, journalId=1302319053441957962, language=EN, title=Heterologous expression of Panax notoginseng tonoplast intrinsic protein gene PnTIP1;3 in yeast and its role in arsenic transport, columnId=null, journalTitle=Chinese Traditional and Herbal Drugs, columnName=null, runingTitle=null, highlight=null, articleAbstract=Objective Based on transcriptome data from roots of Panax notoginseng under arsenic (As) stress, the tonoplast intrinsic protein gene PnTIP1;3 was cloned. Its functional characteristics in As transport were investigated through bioinformatic analysis and heterologous expression in yeast. Methods Bioinformatic tools were used to analyze the sequence features and promoter cis-acting elements of PnTIP1;3. A yeast heterologous expression system was constructed, and the role of PnTIP1;3 in cellular As transport was systematically evaluated through phenotypic assays, intracellular As content and subcellular distribution measurement by ICP-MS, and analysis of relevant antioxidant indicators. Results The tonoplast intrinsic protein gene PnTIP1;3 was cloned, with an open reading frame of 750 bp encoding 250 amino acids. Phylogenetic analysis indicated that it belongs to the TIP1 subfamily. Promoter analysis revealed the presence of multiple cis-acting elements related to hormones, growth and development, and stress responses. As tolerance assays showed that the half-maximal effective concentration (EC50) of yeast transformants heterologously expressing PnTIP1;3 (TPnTIP1;3) was 5.36 mmol/L, higher than that of the empty vector control (5.03 mmol/L). As treatment inhibited the growth of both strains, but the empty vector control was more severely affected. ICP-MS analysis indicated that total As accumulation was significantly higher in TPnTIP1;3 than in the control, and the subcellular distribution of As followed the pattern: cell wall > vacuole > cytoplasm. Under As stress, reactive oxygen species (ROS) levels were significantly decreased (P < 0.001) in TPnTIP1;3, while the activities of superoxide dismutase (SOD) and catalase (CAT), as well as the contents of glutathione (GSH) and metallothionein (MT), were significantly increased (P < 0.05). Correlation analysis further demonstrated significant positive relationships between these antioxidant indicators and both total As content and cell wall As concentration. Conclusion The P. notoginseng tonoplast intrinsic protein gene PnTIP1;3, a member of the TIP1 subfamily, enhances yeast tolerance to As stress by promoting As accumulation and vacuolar sequestration, and through activation of the host antioxidant system., authors=ZHENG Taixiong, ZHANG Xingkai, TANG Shushuang, WEI Fugang, WANG Yanlin, LONG Xiannv, CAO Guanhua, HE Sen, authorsList=ZHENG Taixiong, ZHANG Xingkai, TANG Shushuang, WEI Fugang, WANG Yanlin, LONG Xiannv, CAO Guanhua, HE Sen, authorCompany=null, correspAuthors=null, authorNote=null, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=null, pdfFileSize=null, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=null), CN=ArticleExt(id=1304388058552496714, articleId=1304388058355364425, tenantId=1146029695717560320, journalId=1302319053441957962, language=CN, title=三七液泡膜内在蛋白基因PnTIP1;3的酵母异源表达及其砷转运功能解析, columnId=1304140203967410414, journalTitle=中草药, columnName=药材与资源, runingTitle=null, highlight=null, articleAbstract=目的 基于砷(As)胁迫下三七Panax notoginseng根的转录组数据,克隆液泡膜内在蛋白基因PnTIP1;3,通过生物信息学分析和酵母异源表达实验,解析其在As转运过程中的功能特性。方法 利用生物信息学工具分析基因PnTIP1;3的序列特征及其启动子区顺式作用元件;构建酵母异源表达体系,通过表型实验、ICP-MS测定细胞内As含量与亚细胞分布,并检测相关抗氧化指标,系统评估PnTIP1;3在细胞As转运中的功能。结果 克隆获得三七液泡膜内在蛋白基因PnTIP1;3,其开放阅读框为750 bp,编码250个氨基酸,系统进化分析表明其属于TIP1亚族。启动子分析显示其含有多个激素、生长发育及胁迫响应相关的顺式作用元件。As耐性实验表明,异源表达PnTIP1;3的酵母转化子(TPnTIP1;3)的EC₅₀为5.36 mmol/L,略高于空载对照(5.03 mmol/L)。As处理对2种菌株生长均产生抑制,但对空载菌株的抑制更为显著。ICP-MS分析显示,TPnTIP1;3菌株中总As积累量显著高于对照,亚细胞组分中As含量表现为细胞壁>液泡>细胞质。在As胁迫下,TPnTIP1;3中活性氧(reactive oxygen species,ROS)水平显著降低(P<0.001),而超氧化物歧化酶(superoxide dismutase,SOD)、过氧化氢酶(catalase,CAT)活性以及谷胱甘肽(glutathione,GSH)和金属硫蛋白(metallothionein,MT)含量均显著上升(P<0.05)。相关性分析进一步表明,这些抗氧化指标与总As含量及细胞壁As浓度呈显著正相关。结论 三七液泡膜内在蛋白基因PnTIP1;3属于TIP1亚族,其异源表达可促进酵母对As的积累与液泡区隔化储存,并通过激活宿主抗氧化系统增强对As胁迫的耐受能力。, authors=郑泰雄1,2,3,4, 张兴开1, 唐树爽1, 魏富刚5, 王艳林5, 龙仙女1, 曹冠华1,2,3,4, 贺森1,2,3,4, authorsList=郑泰雄, 张兴开, 唐树爽, 魏富刚, 王艳林, 龙仙女, 曹冠华, 贺森, authorCompany=1 云南中医药大学中药学院, 云南 昆明 650500; 2 云南中医药大学中药学院暨云南省南药可持续利用重点实验室, 云南 昆明 650500; 3 道地药材品质保障与资源持续利用全国重点实验室, 北京 100700; 4 云南中医药大学 中药研究国际科技合作基地, 云南 昆明 650500; 5 文山苗乡三七科技有限公司, 云南 文山 663108, correspAuthors=曹冠华, authorNote=郑泰雄: 郑泰雄,硕士研究生,研究方向为中药资源开发。E-mail:2558707769@qq.com, correspAuthorsNote=null, copyrightStatement=null, copyrightOwner=null, extLink=null, articleAbsUrl=null, sourceXml=null, magXml=null, pdfUrl=null, pdf=dSWnuT5e2SN6XtuDiQIW5w==, pdfFileSize=2107953, pdfExtLink=null, richHtmlUrl=null, mobilePdfUrl=null, reviewReport=null, pdfFirstPage=null, abstractGraph=null, abstractGraphContent=null, abstractVideo=null, citation=null, cebUrl=null, magXmlContent=null, mapNumber=null, fund=云南省基础研究计划重点项目 (202501AS070141); 国家自然科学基金项目 (82260743); 国家自然科学基金项目 (82360750); 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Determination of the authenticity and origin of Panax notoginseng: A review [J]. J Aoac International, 2022, 105(6): 1708-1718. 张丹, 冯丽颖, 高立志. 三七生物资源研究与利用进展[J]. 生物资源, 2020, 42(1): 61-66. 吴涵, 何忠俊, 孟溪, 等. 三七-土壤系统重金属分布特征及健康风险评价[J]. 农业环境科学学报, 2023, 42(7): 1477-1486. 陶亮, 包立, 刘源, 等. 云南不同产地三七的重金属吸收累积特征研究[J]. 中国农学通报, 2018, 34(34): 74-81. Garbinski L D, Rosen B P, Chen J. Pathways of arsenic uptake and efflux [J]. Environ Int, 2019, 126: 585-597. Li J J, Yang L, Miao C P, et al. Impact of rhizosphere microorganisms on arsenic (As) transformation and accumulation in a traditional Chinese medical plant [J]. Environ Sci Pollut Res, 2021, 28(43): 60923-60934. 李泽东. 不同磷水平处理对三七砷吸收的影响及调控机制研究[D]. 昆明: 云南中医药大学, 2019. Zeng X C, Jiang Y X, Fan X T, et al. Effects of sulfate application on inhibiting accumulation and alleviating toxicity of arsenic in Panax notoginseng grown in arsenic-polluted soil [J]. Water Air Soil Pollut, 2016, 227(5): 148. Deng F L, Liu X, Chen Y S, et al. Aquaporins mediated arsenite transport in plants: Molecular mechanisms and applications in crop improvement [J]. Crit Rev Environ Sci Technol, 2020, 50(16): 1613-1639. Xi Y M, Han B L, Kong F T, et al. Enhancement of arsenic uptake and accumulation in green microalga Chlamydomonas reinhardtii through heterologous expression of the phosphate transporter DsPht1[J]. J Hazard Mater, 2023, 459: 132130. Kumar K, Mosa K A, Meselhy A G, et al. Molecular insights into the plasma membrane intrinsic proteins roles for abiotic stress and metalloids tolerance and transport in plants [J]. Indian J Plant Physiol, 2018, 23(4): 721-730. Hussain A, Tanveer R, Mustafa G, et al. Comparative phylogenetic analysis of aquaporins provides insight into the gene family expansion and evolution in plants and their role in drought tolerant and susceptible chickpea cultivars [J]. Genomics, 2020, 112(1): 263-275. Salehi M. Epichloe endophyte modifies antioxidative defense and aquaporin genes expression in response to Ni contamination in Lolium perenne [J]. J Plant Process Funct, 2021, 10(43): 55-63. Zhang L D, Song L Y, Dai M J, et al. Cadmium promotes the absorption of ammonium in hyperaccumulator Solanum nigrum L. mediated by ammonium transporters and aquaporins [J]. Chemosphere, 2022, 307(Pt 3): 136031. Ariani A, Barozzi F, Sebastiani L, et al. AQUA1 is a mercury sensitive poplar aquaporin regulated at transcriptional and post-translational levels by Zn stress [J]. Plant Physiol Biochem, 2019, 135: 588-600. He Z Y, Yan H L, Chen Y S, et al. An aquaporin PvTIP41 from Pteris vittata may mediate arsenite uptake [J]. New Phytol, 2016, 209(2): 746-761. 曹冠华, 张兴开, 柏旭, 等. 三七细胞色素P450酶基因PnCyp450_3响应丛枝菌根真菌诱导的表达特性分析[J]. 中草药, 2022, 53(21): 6848-6856. Zhang W J, Zhou Y W, Zhang Y, et al. Protein phosphorylation: A molecular switch in plant signaling [J]. Cell Rep, 2023, 42(7): 112729. 王海波, 吴贞莹, 郭俊云. 小桐子植物特异性酪蛋白激酶PS-CK1-5基因的克隆及原核表达分析[J]. 热带亚热带植物学报, 2024, 32(1): 93-100. 冯燕, 吴志会, 曹雨欣, 等. 小麦酪蛋白激酶TaCK2α启动子的克隆与启动活性分析[J]. 河北农业大学学报, 2025, 48(2): 54-61. Zhu J, Wang W S, Yan D W, et al. CK2 promotes jasmonic acid signaling response by phosphorylating MYC2 in Arabidopsis [J]. Nucleic Acids Res, 2023, 51(2): 619-630. 贲琳丽. 水稻水通道蛋白TIP和PIP家族对非生物胁迫的响应及OsTIP12的表达特征[D]. 扬州: 扬州大学, 2024. Khan I, Awan S A, Rizwan M, et al. Arsenic behavior in soil-plant system and its detoxification mechanisms in plants: A review [J]. Environ Pollut, 2021, 286: 117389. 柯汉玲, 祖艳群. 三年生三七生长、光合特征及砷含量对土壤砷胁迫的响应[J]. 云南农业大学学报: 自然科学, 2016, 31(6): 1065-1072. Lu T Q, Wang X N, Cui X L, et al. Physiological and metabolomic analyses reveal that Fe3O4 nanoparticles ameliorate cadmium and arsenic toxicity in Panax notoginseng [J]. Environ Pollut, 2023, 337: 122578. Cao G H, Bai X, Zhang C R, et al. Physiological response and transcriptome profiling reveal phosphate-mediated amelioration of arsenic accumulation and toxicity in Panax notoginseng [J]. Environ Exp Bot, 2023, 206: 105136. Handa N, Gupta P, Khanna K, et al. Aquaporin-mediated transport: Insights into metalloid trafficking [J]. Physiol Plant, 2022, 174(3): e13687. Feng Z J, Liu N, Zhang G W, et al. Investigation of the AQP family in soybean and the promoter activity of TIP26 in heat stress and hormone responses [J]. Int J Mol Sci, 2019, 20(2): 262. Rodrigues M I, Bravo J P, Sassaki F T, et al. The tonoplast intrinsic aquaporin (TIP) subfamily of Eucalyptus grandis: Characterization of EgTIP2, a root-specific and osmotic stress-responsive gene [J]. Plant Sci, 2013, 213: 106-113. Kurowska M M, Wiecha K, Gajek K, et al. Drought stress and re-watering affect the abundance of TIP aquaporin transcripts in barley [J]. PLoS One, 2019, 14(12): e0226423. 陈静, 赵玉全, 黄锡金, 等. 外源脱落酸对甘蓝型油菜镉胁迫的生理响应及基因表达分析[J]. 植物遗传资源学报, 2024, 25(12): 2107-2121. 熊明彪, 饶逸驰, 王乾鑫, 等. 铅、镉胁迫下蜀葵喷施赤霉素与油菜间作对油菜重金属积累的影响[J]. 北方园艺, 2022(17): 17-25. Karle S B, Kumar K. Rice tonoplast intrinsic protein member OsTIP12 confers tolerance to arsenite stress [J]. J Hazard Mater, 2024, 465: 133078. Wang Y Q, Kang Y, Yu W C, et al. AtTIP22 facilitates resistance to zinc toxicity via promoting zinc immobilization in the root and limiting root-to-shoot zinc translocation in Arabidopsis thaliana [J]. Ecotoxicol Environ Saf, 2022, 233: 113333. Zhang W Y, Yang S H, Feng Y M, et al. The tonoplast-localized OsTIP21 is involved in aluminum detoxification in rice [J]. Plant Physiol Biochem, 2024, 215: 109063. Sun H Y, Li L C, Lou Y F, et al. The bamboo aquaporin gene PeTIP41–1 confers drought and salinity tolerance in transgenic Arabidopsis [J]. Plant Cell Rep, 2017, 36(4): 597-609. Wu Z C, Liu S, Zhao J, et al. Comparative responses to silicon and selenium in relation to antioxidant enzyme system and the glutathione-ascorbate cycle in flowering Chinese cabbage (Brassicacampestris L. ssp. chinensis var. utilis) under cadmium stress [J]. Environ Exp Bot, 2017, 133: 1-11. Kim Y O, Gwon Y, Kim J. Exogenous cysteine improves mercury uptake and tolerance in Arabidopsis by regulating the expression of heavy metal chelators and antioxidative enzymes [J]. Front Plant Sci, 2022, 13: 898247. Pang Y Q, Li L J, Ren F, et al. Overexpression of the tonoplast aquaporin AtTIP51 conferred tolerance to boron toxicity in Arabidopsis [J]. J Genet Genomics, 2010, 37(6): 389-397.)
Objective Based on transcriptome data from roots of Panax notoginseng under arsenic (As) stress, the tonoplast intrinsic protein gene PnTIP1;3 was cloned. Its functional characteristics in As transport were investigated through bioinformatic analysis and heterologous expression in yeast. Methods Bioinformatic tools were used to analyze the sequence features and promoter cis-acting elements of PnTIP1;3. A yeast heterologous expression system was constructed, and the role of PnTIP1;3 in cellular As transport was systematically evaluated through phenotypic assays, intracellular As content and subcellular distribution measurement by ICP-MS, and analysis of relevant antioxidant indicators. Results The tonoplast intrinsic protein gene PnTIP1;3 was cloned, with an open reading frame of 750 bp encoding 250 amino acids. Phylogenetic analysis indicated that it belongs to the TIP1 subfamily. Promoter analysis revealed the presence of multiple cis-acting elements related to hormones, growth and development, and stress responses. As tolerance assays showed that the half-maximal effective concentration (EC50) of yeast transformants heterologously expressing PnTIP1;3 (TPnTIP1;3) was 5.36 mmol/L, higher than that of the empty vector control (5.03 mmol/L). As treatment inhibited the growth of both strains, but the empty vector control was more severely affected. ICP-MS analysis indicated that total As accumulation was significantly higher in TPnTIP1;3 than in the control, and the subcellular distribution of As followed the pattern: cell wall > vacuole > cytoplasm. Under As stress, reactive oxygen species (ROS) levels were significantly decreased (P < 0.001) in TPnTIP1;3, while the activities of superoxide dismutase (SOD) and catalase (CAT), as well as the contents of glutathione (GSH) and metallothionein (MT), were significantly increased (P < 0.05). Correlation analysis further demonstrated significant positive relationships between these antioxidant indicators and both total As content and cell wall As concentration. Conclusion The P. notoginseng tonoplast intrinsic protein gene PnTIP1;3, a member of the TIP1 subfamily, enhances yeast tolerance to As stress by promoting As accumulation and vacuolar sequestration, and through activation of the host antioxidant system.
ZHENG Taixiong, ZHANG Xingkai, TANG Shushuang, WEI Fugang, WANG Yanlin, LONG Xiannv, CAO Guanhua, HE Sen.
Heterologous expression of Panax notoginseng tonoplast intrinsic protein gene PnTIP1;3 in yeast and its role in arsenic transport[J].
Chinese Traditional and Herbal Drugs,
2026
, 57
(10)
: 3926
-3939
.
DOI: 10.7501/j.issn.0253-2670.2026.10.021
Ji C, Zhang Q, Shi R, et al. Determination of the authenticity and origin of Panax notoginseng: A review [J]. J Aoac International, 2022, 105(6): 1708-1718. 张丹, 冯丽颖, 高立志. 三七生物资源研究与利用进展[J]. 生物资源, 2020, 42(1): 61-66. 吴涵, 何忠俊, 孟溪, 等. 三七-土壤系统重金属分布特征及健康风险评价[J]. 农业环境科学学报, 2023, 42(7): 1477-1486. 陶亮, 包立, 刘源, 等. 云南不同产地三七的重金属吸收累积特征研究[J]. 中国农学通报, 2018, 34(34): 74-81. Garbinski L D, Rosen B P, Chen J. Pathways of arsenic uptake and efflux [J]. Environ Int, 2019, 126: 585-597. Li J J, Yang L, Miao C P, et al. Impact of rhizosphere microorganisms on arsenic (As) transformation and accumulation in a traditional Chinese medical plant [J]. Environ Sci Pollut Res, 2021, 28(43): 60923-60934. 李泽东. 不同磷水平处理对三七砷吸收的影响及调控机制研究[D]. 昆明: 云南中医药大学, 2019. Zeng X C, Jiang Y X, Fan X T, et al. Effects of sulfate application on inhibiting accumulation and alleviating toxicity of arsenic in Panax notoginseng grown in arsenic-polluted soil [J]. Water Air Soil Pollut, 2016, 227(5): 148. Deng F L, Liu X, Chen Y S, et al. Aquaporins mediated arsenite transport in plants: Molecular mechanisms and applications in crop improvement [J]. Crit Rev Environ Sci Technol, 2020, 50(16): 1613-1639. Xi Y M, Han B L, Kong F T, et al. Enhancement of arsenic uptake and accumulation in green microalga Chlamydomonas reinhardtii through heterologous expression of the phosphate transporter DsPht1[J]. J Hazard Mater, 2023, 459: 132130. Kumar K, Mosa K A, Meselhy A G, et al. Molecular insights into the plasma membrane intrinsic proteins roles for abiotic stress and metalloids tolerance and transport in plants [J]. Indian J Plant Physiol, 2018, 23(4): 721-730. Hussain A, Tanveer R, Mustafa G, et al. Comparative phylogenetic analysis of aquaporins provides insight into the gene family expansion and evolution in plants and their role in drought tolerant and susceptible chickpea cultivars [J]. Genomics, 2020, 112(1): 263-275. Salehi M. Epichloe endophyte modifies antioxidative defense and aquaporin genes expression in response to Ni contamination in Lolium perenne [J]. J Plant Process Funct, 2021, 10(43): 55-63. Zhang L D, Song L Y, Dai M J, et al. Cadmium promotes the absorption of ammonium in hyperaccumulator Solanum nigrum L. mediated by ammonium transporters and aquaporins [J]. Chemosphere, 2022, 307(Pt 3): 136031. Ariani A, Barozzi F, Sebastiani L, et al. AQUA1 is a mercury sensitive poplar aquaporin regulated at transcriptional and post-translational levels by Zn stress [J]. Plant Physiol Biochem, 2019, 135: 588-600. He Z Y, Yan H L, Chen Y S, et al. An aquaporin PvTIP41 from Pteris vittata may mediate arsenite uptake [J]. New Phytol, 2016, 209(2): 746-761. 曹冠华, 张兴开, 柏旭, 等. 三七细胞色素P450酶基因PnCyp450_3响应丛枝菌根真菌诱导的表达特性分析[J]. 中草药, 2022, 53(21): 6848-6856. Zhang W J, Zhou Y W, Zhang Y, et al. Protein phosphorylation: A molecular switch in plant signaling [J]. Cell Rep, 2023, 42(7): 112729. 王海波, 吴贞莹, 郭俊云. 小桐子植物特异性酪蛋白激酶PS-CK1-5基因的克隆及原核表达分析[J]. 热带亚热带植物学报, 2024, 32(1): 93-100. 冯燕, 吴志会, 曹雨欣, 等. 小麦酪蛋白激酶TaCK2α启动子的克隆与启动活性分析[J]. 河北农业大学学报, 2025, 48(2): 54-61. Zhu J, Wang W S, Yan D W, et al. CK2 promotes jasmonic acid signaling response by phosphorylating MYC2 in Arabidopsis [J]. Nucleic Acids Res, 2023, 51(2): 619-630. 贲琳丽. 水稻水通道蛋白TIP和PIP家族对非生物胁迫的响应及OsTIP12的表达特征[D]. 扬州: 扬州大学, 2024. Khan I, Awan S A, Rizwan M, et al. Arsenic behavior in soil-plant system and its detoxification mechanisms in plants: A review [J]. Environ Pollut, 2021, 286: 117389. 柯汉玲, 祖艳群. 三年生三七生长、光合特征及砷含量对土壤砷胁迫的响应[J]. 云南农业大学学报: 自然科学, 2016, 31(6): 1065-1072. Lu T Q, Wang X N, Cui X L, et al. Physiological and metabolomic analyses reveal that Fe3O4 nanoparticles ameliorate cadmium and arsenic toxicity in Panax notoginseng [J]. Environ Pollut, 2023, 337: 122578. Cao G H, Bai X, Zhang C R, et al. Physiological response and transcriptome profiling reveal phosphate-mediated amelioration of arsenic accumulation and toxicity in Panax notoginseng [J]. Environ Exp Bot, 2023, 206: 105136. Handa N, Gupta P, Khanna K, et al. Aquaporin-mediated transport: Insights into metalloid trafficking [J]. Physiol Plant, 2022, 174(3): e13687. Feng Z J, Liu N, Zhang G W, et al. Investigation of the AQP family in soybean and the promoter activity of TIP26 in heat stress and hormone responses [J]. Int J Mol Sci, 2019, 20(2): 262. Rodrigues M I, Bravo J P, Sassaki F T, et al. The tonoplast intrinsic aquaporin (TIP) subfamily of Eucalyptus grandis: Characterization of EgTIP2, a root-specific and osmotic stress-responsive gene [J]. Plant Sci, 2013, 213: 106-113. Kurowska M M, Wiecha K, Gajek K, et al. Drought stress and re-watering affect the abundance of TIP aquaporin transcripts in barley [J]. PLoS One, 2019, 14(12): e0226423. 陈静, 赵玉全, 黄锡金, 等. 外源脱落酸对甘蓝型油菜镉胁迫的生理响应及基因表达分析[J]. 植物遗传资源学报, 2024, 25(12): 2107-2121. 熊明彪, 饶逸驰, 王乾鑫, 等. 铅、镉胁迫下蜀葵喷施赤霉素与油菜间作对油菜重金属积累的影响[J]. 北方园艺, 2022(17): 17-25. Karle S B, Kumar K. Rice tonoplast intrinsic protein member OsTIP12 confers tolerance to arsenite stress [J]. J Hazard Mater, 2024, 465: 133078. Wang Y Q, Kang Y, Yu W C, et al. AtTIP22 facilitates resistance to zinc toxicity via promoting zinc immobilization in the root and limiting root-to-shoot zinc translocation in Arabidopsis thaliana [J]. Ecotoxicol Environ Saf, 2022, 233: 113333. Zhang W Y, Yang S H, Feng Y M, et al. The tonoplast-localized OsTIP21 is involved in aluminum detoxification in rice [J]. Plant Physiol Biochem, 2024, 215: 109063. Sun H Y, Li L C, Lou Y F, et al. The bamboo aquaporin gene PeTIP41–1 confers drought and salinity tolerance in transgenic Arabidopsis [J]. Plant Cell Rep, 2017, 36(4): 597-609. Wu Z C, Liu S, Zhao J, et al. Comparative responses to silicon and selenium in relation to antioxidant enzyme system and the glutathione-ascorbate cycle in flowering Chinese cabbage (Brassicacampestris L. ssp. chinensis var. utilis) under cadmium stress [J]. Environ Exp Bot, 2017, 133: 1-11. Kim Y O, Gwon Y, Kim J. Exogenous cysteine improves mercury uptake and tolerance in Arabidopsis by regulating the expression of heavy metal chelators and antioxidative enzymes [J]. Front Plant Sci, 2022, 13: 898247. Pang Y Q, Li L J, Ren F, et al. Overexpression of the tonoplast aquaporin AtTIP51 conferred tolerance to boron toxicity in Arabidopsis [J]. J Genet Genomics, 2010, 37(6): 389-397.